Zootaxa 3905 (2): 193–208 ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Article ZOOTAXA Copyright © 2015 Magnolia Press ISSN 1175-5334 (online edition) http://dx.doi.org/10.11646/zootaxa.3905.2.2 http://zoobank.org/urn:lsid:zoobank.org:pub:3C2F37A5-262F-4C36-BF66-FD3254E2F147 Two new marine Gastrotricha from the Indian Ocean coast of South Africa

M. ANTONIO TODARO1,4, RENZO PERISSINOTTO2,3 & SARAH J. BOWNES3 1Department of Life Sciences, University of Modena e Reggio Emilia, via Campi, 231/D, I-41125 Modena, Italy 2DST/NRF South African Research Chair in Shallow Water Ecosystems, Nelson Mandela Metropolitan University, P.O. Box 77000, Port Elizabeth 6031, South Africa 3School of Life Sciences, University of KwaZulu-Natal, Westville Campus, Private Bag X54001, Durban 4000, South Africa 4Corresponding author. E-mail: [email protected]

Abstract

The study is part of a larger research programme aimed at shedding light on the communities of the subtropical east coast province of KwaZulu-Natal, South Africa. In previous papers, faunistic and preliminary taxonomic data on ma- rine and freshwater gastrotrichs found in the iSimangaliso Wetland Park, were reported. Here two new interesting marine macrodasyidan species in the families Dactylopodolidae and are described based on observations carried out on living specimens and using differential interference contrast microscopy. The two novel species are named in honor of two great South African icons recently deceased: Nadine Gordimer and Nelson Mandela. Dactylopodola na- dine sp. n. is the third species in the to bear red eye-spots; it can easily be distinguished from the closely-related red-eyed D. baltica and D. roscovita by its smaller size (Total length = 230 μm vs 275 μm vs 450 μm, respectively) and the lower number of adhesive tubes of the anterior, lateral and posterior series (on each side: 3, 4 and 4 vs 5, 6 and 8 vs 2, 9 and 12–15). Pseudostomella mandela sp. n. is a fairly large species (up to 481 μm in length), with a cuticular covering made up of tetrancres and relatively long caudal pedicles (up to 44 μm in length ). The most evident autoapomorphic trait of the new species is the presence of 7 pairs of ‘cirrata’ tubes, two emerging in a lateral position along the pharyngeal region and five from the dorsolateral sides of the trunk. Additional relevant taxonomic characters include: 4 tubes of the anterior series, 11 tubes of the ventrolateral series and 3 tubes of the posterior series per side, 5 papillae on the dorsal mar- gin and 6 papillae on the ventral margin of the oral palps. The high number of putative new species discovered among the South African gastrotrich fauna during our relatively short survey, highlights the relevance of this region with regard to the diversity of this group and stresses once again the importance of investigating new geographic areas in order to im- prove our understanding of global gastrotrich biodiversity and species richness.

Key words: biodiversity, Gastrotrichs, Meiofauna, KwaZulu-Natal, South Africa, , new species

Introduction

The Phylum Gastrotricha includes meiofaunal-sized, vermiform invertebrates found in both freshwater and marine ecosystems (e.g., Balsamo & Todaro 2002; Todaro & Hummon 2008). The group comprises about 850 species divided into the two orders, and Macrodasyida (Balsamo et al. 2009; Hummon & Todaro 2010; Todaro 2014). The Chaetonotida includes taxa found in marine, brackish or freshwater habitats, whereas Macrodasyida generally includes species living interstitially in the sand of marine ecosystems (but see e.g., Todaro et al. 2012). The systematics and alpha biodiversity of the entire Phylum are in a state of flux as shown in the recent in-group taxa reassessment based on new phylogenetic data (e.g. Todaro et al. 2011a, 2012a, 2014; Kanneby et al. 2013; Guidi et al. 2014; Todaro et al. 2014) and the continual description of new species (e.g. freshwater: Kånneby 2013; Kolicka et al. 2013; Suzuki et al. 2013; Schwank & Kanneby 2014; marine: Hochberg et al. 2013; Kieneke et al. 2013; Lee et al. 2013; Todaro 2013; Todaro & Leasi 2013; Araujo et al. 2014; Atherton 2014; Gilsa et al. 2014; Hochberg et al. 2014; Kånneby et al. 2014; Lee & Chang 2014; Lee et al. 2014). In terms of biodiversity, meiofauna and small macrobenthos are still relatively poorly known from South Africa. Although some progress has been made in recent years (e.g., Hendricks & Gibbons 2010; Todaro et al.

Accepted by W. Sterrer: 4 Dec. 2014; published: 12 Jan. 2015 193 Taxonomic affinities. Within Thaumastodermatinae, the genera Tetranchyroderma and Pseudostomella, the number of prongs forming the peculiar scales called ancres, which composes the body cuticular armature, has been regarded as the single most useful taxonomic trait to classify species (e.g., Lee & Chang 2002; Todaro 2002; but see Todaro et al. 2011a). Consequently, in the genus Pseudostomella three basic species groups are envisaged based on the type of pronged spines i.e., species characterized by triancres (3 prongs), tetrancres (4 prongs) or pentancres (5 prongs). Based on the type of ancres, Pseudostomella mandela n. sp. approaches six other species all characterized by a tetrancrous covering: P. andamanica Rao, 1993, P. indica Rao, 1970, P. koreana Lee & Chang, 2002, P. longifurca Lee & Chang, 2002, P. m al ayi ca Renaud-Mornant, 1967 and P. roscovi ta Swedmark, 1956 (see Todaro 2012). However, based on the number of dorsal papillae (5) on the preoral palps, the new species is most similar to P. longifurca and P. indica. The number and distribution of the adhesive tubes is useful in discriminating the three taxa e.g., P. longifurca bears 5 TbA and 7 TbP per side and P. indi ca 2 TbA and 5 TbP, in contrast with the new species that exhibits 4 TbA and 4 TbP per side. P. mandela n. sp. is further distinguished as the only species in the genus that possesses cirrata tubes (7 pairs). The finding of an additional new taxon seems to support the idea that Pseudostomella species appear to have a relatively restricted geographic range, at least compared to the wide distribution and cosmopolitan nature of many other gastrotrichs (cf. Todaro et al. 1996; Artois et al. 2011; Curini-Galletti et al. 2012; Kånneby et al. 2012; Kieneke et al. 2012).

Conclusive remarks

Prior to our study, South Africa was ‘terra incognita’ with respect to the gastrotrich fauna; overall we have sampled only 5–6 marine localities and 3–4 freshwater ponds, all in the Province of KwaZulu-Natal, from where we have so far censused about 30 marine and 15 freshwater putative species. Of these, 4 marine and 1 freshwater species have already been described as new to science (Todaro et al., 2011b, 2013 and present study) and many others appear to bear traits that would classify them as new taxa. Although estimates about the uniqueness of the South African gastrotrich fauna (at least of KwaZulu-Natal) can only be made at the end of the ongoing taxonomic investigation, the high incidence of potential new species among the fauna recorded during our relatively short survey, makes immediately manifest the relevance of this region with regard to the diversity of this group. This certainly calls for a widening of studies to other South African provinces, as the diversity of climatic, sedimentary and orographic features within this area is remarkable and is generally reflected in its rich biodiversity. In a larger framework, results of this initial investigation stress once again the importance of investigations carried out in new geographic areas for the improvement of our understanding of the overall species richness and global gastrotrich biodiversity (cf. Todaro in Appeltans et al. 2012; Todaro et al. 2014).

Acknowledgements

The staff and management at iSimangaliso Wetlands Park Authority and Ezemvelo KZN Wildlife are thanked for their support and co-operation. We thank the Director, Nelisha Murugan, and the Principal Technician, Phillip Christopher, of the Microscopy and Micro-analysis Unit of the University of KwaZulu-Natal for kindly providing facilities and assistance towards the success of this study. We are also very grateful to Nelson Miranda, Nicola Carrasco and David Dyer for their invaluable assistance with field collections. Funding for this study was provided by the National Research Foundation (NRF, Pretoria), the University of KwaZulu-Natal (UKZN, Durban) and the University of Modena and Reggio Emilia, TIOME project. The study falls within the framework of the strategic research line—Environment, food and health (WP2)—of UNIMORE.

References

Appeltans, W., Ahyong, S.T., Anderson, G., Angel, M.V., Artois, T., Bailly, N., Bamber, R., Barber, A., Bartsch, I., Berta, A., Blazewicz-Paszkowycz, M., Bock, P., Boxshall, G., Boyko, C.B., Nunes Brandão, S., Bray, R.A., Bruce, N.L., Cairns, S.D., Chan, T-Y., Cheng, L., Collins, A.G., Cribb, T., Curini-Galletti, M., Dahdouh-Guebas, F., Davie, P.J.F., Dawson,

TWO NEW MARINE GASTROTRICHA FROM SOUTH AFRICA Zootaxa 3905 (2) © 2015 Magnolia Press · 205 M.N., De Clerck, O., Decock, W., De Grave, S., de Voogd, N.J., Domning, D.P., Emig, C.C., Erséus, C., Eschmeyer, W., Fauchald, K., Fautin, D.F., Feist, S.W., Fransen, C.H.J.M , Furuya, H., Garcia-Alvarez, O., Gerken, S., Gibson, D., Gittenberger, A., Gofas, S., Gómez-Daglio, L., Gordon, D.P., Guiry, M.D., Hernandez, F., Hoeksema, B.W., Hopcroft, R.R., Jaume, D., Kirk, P., Koedam, N., Koenemann, S., Kolb, J.B., Kristensen, R.M., Kroh, A., Lambert, G., Lazarus, D.B., Lemaitre, R., Longshaw, M., Lowry, J., Macpherson, E., Madin, L.P., Mah, C., Mapstone, G., McLaughlin, P.A., Meland, J.M.K., Messing, C.G., Mills, C.E., Molodtsova, T.N., Mooi, R., Neuhaus, B., Ng, P.K.L., Nielsen, C., Norenburg, J., Opresko, D.M., Osawa, M., Paulay, G., Perrin, W., Pilger, J.F., Poore, G.C.B., Pugh, P., Read, G.B., Reimer, J.D., Rius, M., Rocha, R.M., Saiz-Salinas, J.I., Scarabino, V., Schierwater, B., Schmidt-Rhaesa, A., Schnabel, K.E., Schotte, M., Schuchert, P., Schwabe, E., Segers, H., Self-Sullivan, C., Shenkar, N., Siegel, V., Sterrer, W., Stöhr, S., Swalla, B., Tasker, M.L., Thuesen, E.V., Timm, T., Todaro, M.A., Turon, X., Tyler, S., Uetz, P., van der Land, J., Vanhoorne, B., van Ofwegen, L.P., van Soest, R.W.M., Vanaverbeke, J., Walker-Smith, G., Walter, T.C., Warren, A., Williams, G.C., Wilson, S.P. & Costello, M.J. (2012) The Magnitude of Global Marine Species Diversity. Current Biology, 22, 2189–2202. http://dx.doi.org/10.1016/j.cub.2012.09.036 Araujo, T.Q., Balsamo, M. & Garraffoni, A.R.S. (2014) A new species of Pseudostomodella (Gastrotricha, Thaumastodermatidae) from Brazil. Marine Biodiversity, 44, 243–248. http://dx.doi.org/10.1007/s12526-014-0263-y Artois, T., Fontaneto, D., Hummon, W.D., Mcinnes, S.J., Todaro, M.A., Sørensen, M.V. & Zullini, A. (2011) Ubiquity of microscopic ? Evidence from the morphological approach in species identification. In: Fontaneto, D. (Ed.), Biogeography of microscopic organisms: is everything small everywhere? Cambridge University Press, Cambridge, pp. 244–283. http://dx.doi.org/10.1017/CBO9780511974878.014 Artois, T., Willems, W., Reygel, P. & Schockaert, E. (2013) Brachyrhynchus n. gen. n. sp., a new genus of Polycystididae Graff, 1905 (Rhabdocoela: Kalyptorhynchia), with the description of three new species from the Mediterranean and the Indian Ocean. Zootaxa, 3635 (2), 127–136. http://dx.doi.org/10.11646/zootaxa.3635.2.3 Atherton, S. (2014) Urodasys poculostylis sp. nov., a new stylet-bearing gastrotrich (Macrodasyida) from Capron Shoal, Florida. Marine Biology Research, 5, 530–536. http://dx.doi.org/10.1080/17451000.2013.831179 Balsamo, M., d'Hondt, J.L., Pierboni, L. & Grilli, P. (2009) Taxonomic and nomenclatural notes on freshwater Gastrotricha. Zootaxa, 2158, 1–19. Balsamo, M., Todaro, M.A. (2002) Gastrotricha. In: Rundle, S.D., Robertson, A.I., Schmid-Araya, J.M. (Eds.), Freshwater Meiofauna Biology and Ecology. Backhuys Publisher, Leiden, pp. 45–61. Bownes, S.J. & Perissinotto, R. (2012) Community structure and composition of meiofauna over a sea-induced mouth- breaching event in St Lucia Estuary, South Africa. Marine Ecology Progress Series, 463, 105–126. http://dx.doi.org/10.3354/meps09870 Curini-Galletti, M., Artois, T., Delogu, V., De Smet, W.H., Fontaneto, D., Jondelius, U., Leasi, F., Martinez, A., Meyer- Wachsmuth, I., Nilsson, K.S., Tongiorgi, P., Worsaae, K. & Todaro, M.A. (2012) Patterns of Diversity in Soft-Bodied Meiofauna: Dispersal Ability and Body Size Matter. PLoS ONE, 7 (3), e33801. http://dx.doi.org/10.1371/journal.pone.0033801 Daly, M., Perissinotto, R., Laird, M., Dyer, D. & Todaro, M.A. (2012) Description and ecology of a new species of Edwardsia de Quatrefages, 1842 (Anthozoa, Actiniaria) from the St Lucia Estuary, South Africa. Marine Biology Research, 8, 233–245. http://dx.doi.org/10.1080/17451000.2011.617757 Folk, R.L. & Ward, W.C. (1957) Brazos River Bar: A study in the significance of grain size parameters. Journal of Sediment Petrology, 27, 3–26. Gilsa, A., Kieneke, A., Hochberg, R. & Schmidt-Rhaesa, A. (2014) Two new species of the genus Dactylopodola (Gastrotricha: Macrodasyida) from the Bahamas, with an updated key to the genus. Cahiers De Biologie Marine, 55, 333–345. Gómez, S., Carrasco, N.K. & Morales-Serna, F.N. (2012) A new species of Nitocra Boeck, 1865 (Harpacticoida, Ameiridae, Ameirinae) from South Africa, with notes on its ecology and remarks on the status of Nitocra sewelli husmanni Kunz 1976. ZooKeys, 244, 33–58. http://dx.doi.org/10.3897/zookeys.244.2633 Griffiths, C.L, Robinson, T.B., Lange, L. & Mead, A (2010) Marine Biodiversity in South Africa: An Evaluation of Current States of Knowledge. PLoS ONE, 5, e12008. http://dx.doi.org/10.1371/journal.pone.0012008 Guidi, L., Todaro, M.A., Ferraguti, M. & Balsamo, M. (2014) Reproductive system and spermatozoa ultrastructure support the phylogenetic proximity of Megadasys and Crasiella (Gastrotricha, Macrodasyida). Contributions to Zoology, 83, 119–131. Hendricks, M.G.J. & Gibbons, M.J. (2010) Two new nematode species from Saldanha Bay, South Africa: Perepsilonema benguelae sp. nov. and Leptepsilonema saldanhae sp. nov. Desmodorida, Epsilonematidae). Zootaxa, 2504, 20–30. Hochberg, R., Atherton, S. & Gross, V. (2013) A new species of sublittoral marine gastrotrich, Lepidodasys ligni sp. n. (Macrodasyida, Lepidodasyidae), from the Atlantic coast of Florida. ZooKeys, 289, 1–12. http://dx.doi.org/10.3897/zookeys.289.4764 Hochberg, R., Atherton, S. & Kieneke, A. (2014) Marine Gastrotricha of Little Cayman Island with the description of one new

206 · Zootaxa 3905 (2) © 2015 Magnolia Press TODARO ET AL. species and an initial assessment of meiofaunal diversity. Marine Biodiversity, 44, 89–113. http://dx.doi.org/10.1007/s12526-013-0186-z Hummon, W.D., Balsamo, M. & Todaro, M.A. (1992) Italian marine Gastrotricha: I. Six new and one redescribed species of Chaetonotida. Bollettino di Zoologia, 59, 499–516. http://dx.doi.org/10.1080/11250009209386711 Hummon, W.D. & Todaro, M.A. (2010) Analytic taxonomy and notes on marine, brackish-water and estuarine Gastrotricha. Zootaxa, 2392, 1–32. Hummon, W.D., Todaro, M.A. & Tongiorgi, P. (1993) Italian marine Gastrotricha: II. One new genus and ten new species of Macrodasyida. Bollettino di Zoologia, 60, 109–127. http://dx.doi.org/10.1080/11250009309355798 Kånneby, T. (2013) New species and records of freshwater (Gastrotricha: Chaetonotidae) from Sweden. Zootaxa, 3701 (5), 551–588. http://dx.doi.org/10.11646/zootaxa.3701.5.3 Kånneby, T., Atherton, S. & Hochberg, R. (2014) Two new species of Musellifer (Gastrotricha: Chaetonotida) from Florida and Tobago and the systematic placement of the genus within . Marine Biology Research, 10, 983–995. http://dx.doi.org/10.1080/17451000.2013.872797 Kånneby, T., Todaro, M.A. & Jondelius, U. (2013) Phylogeny of Chaetonotidae and other Paucitubulatina (Gastrotricha: Chaetonotida) and the colonization of aquatic ecosystems. Zoologica Scripta, 42, 88–105. http://dx.doi.org/10.1111/j.1463-6409.2012.00558.x Kånneby, T., Todaro, M.A. & Jondelius, U. (2012) A phylogenetic approach to species delimitation in freshwater Gastrotricha from Sweden. Hydrobiologia, 683, 185–202. http://dx.doi.org/10.1007/s10750-011-0956-1 Kieneke, A., Martinez Arbizu, P. & Fontaneto, D. (2012) Spatially structured populations with a low level of cryptic diversity in European marine Gastrotricha. Molecular Ecology, 21, 1239–1254. http://dx.doi.org/10.1111/j.1365-294X.2011.05421.x Kieneke, A., Narkus, S., Hochberg, R. & Schmidt-Rhaesa, A. (2013) Diplodasys rothei n. sp. (Gastrotricha, Macrodasyida), a new marine gastrotrich species from the Bahamas. Meiofauna Marina, 20, 49–61. Kolicka, M., Kisielewski, J., Nesteruk, T. & Zawierucha, K. (2013) Gastrotricha from the Poznan Palm House—one new subgenus and three new species of freshwater Chaetonotida (Gastrotricha). Zootaxa, 3717 (2), 231–279. http://dx.doi.org/10.11646/zootaxa.3717.2.7 Lee, J.M. & Chang, C.Y. (2002) Pseudostomella gastrotrichs (Macrodasyida, Thaumastodermatidae), form South Korea, with a brief review of the genus. Korean Journal of Biological Sciences, 6, 207–312. http://dx.doi.org/10.2108/zsj.20.481 Lee, J.M. & Chang, C.Y. (2014) A new gastrotrich species of the genus Tetranchyroderma (Macrodasyida: Thaumastodermatidae) from Korea. Proceedings of the Biological Society of Washington, 127, 209–215. http://dx.doi.org/10.2988/0006-324X-127.1.209 Lee, J., Chang, C.Y. & Kim, D. (2014). Dendrodasys duplus, a New Gastrotrich Species (Macrodasyida: Dactylopodolidae) from South Korea. Systematics Evolution and Diversity, 30, 103–107. http://dx.doi.org/10.5635/ASED.2014.30.2.103 Lee, J.M., Kim, D. & Chang, C.Y. (2013) Description of three new Tetranchyroderma gastrotrichs (Macrodasyida: Thaumastodermatidae) from South Korea. Zootaxa, 3709 (5), 483–493. http://dx.doi.org/10.11646/zootaxa.3709.5.6 Pillay, D. & Perissinotto, R. (2009) Community structure of epibenthic meiofauna in the St. Lucia Estuarine Lake (South Africa) during a drought phase. Estuarine Coastal and Shelf Science, 81, 94–104. http://dx.doi.org/10.1016/j.ecss.2008.10.014 Porter, R.N. (2013) South Africa’s first World Heritage Site. In: Perissinotto, R., Stretch, D.D. & Taylor, R.H. (Eds.), Ecology and Conservation of Estuarine Ecosystems, Lake St Lucia as a Global Model. Cambridge University Press, New York, pp. 1–19. Schwank, P. & Kånneby, T. (2014) Contribution to the freshwater gastrotrich fauna of wetland areas of southwestern Ontario (Canada) with redescriptions of seven species and a check-list for North America. Zootaxa, 3811 (4), 463–490. http://dx.doi.org/10.11646/zootaxa.3811.4.3 Suzuki, T.G., Maeda, M. & Furuya, H. (2013) Two new Japanese species of Gastrotricha (Chaetonotida, Chaetonotidae, and Dichaeturidae, Dichaetura), with comments on the diversity of gastrotrichs in rice paddies. Zootaxa, 3691 (2), 229–239. http://dx.doi.org/10.11646/zootaxa.3691.2.3 Todaro, M.A. (2002) An interesting new gastrotrich from littoral meiobenthos (Long Beach Island, USA), with a key to species of Tetranchyroderma (Gastrotricha : Macrodasyida). Journal of the Marine Biological Association of the United Kingdom, 82, 555–563. http://dx.doi.org/10.1017/S0025315402005878 Todaro, M.A. (2012) A new marine gastrotrich from the State of Sao Paulo (Brazil), with a key to species of Pseudostomella (Gastrotricha,Thaumastodermatidae). Zookeys, 223, 39–51.

TWO NEW MARINE GASTROTRICHA FROM SOUTH AFRICA Zootaxa 3905 (2) © 2015 Magnolia Press · 207 http://dx.doi.org/10.3897/zookeys.223.3975 Todaro, M.A. (2013). A new non-naked species of Ptychostomella (Gastrotricha) from Brazil. ZooKeys, 289, 13–24. http://dx.doi.org/10.3897/zookeys.289.4683 Todaro, A. (2014) Gastrotricha. World Register of Marine Species. Available from: http://www.marinespecies.org/ aphia.php?p=taxdetails&id=2078 (accessed 29 July 2014) Todaro, M.A., Kånneby, T., Dal Zotto, M. & Jondelius, U. (2011a) Phylogeny of Thaumastodermatidae (Gastrotricha: Macrodasyida) Inferred from Nuclear and Mitochondrial Sequence Data. PLoS ONE, 6, e17892. http://dx.doi.org/10.1371/journal.pone.0017892 Todaro, M.A., Dal Zotto, M., Bownes, S.J. &Perissinotto, R. (2011b) First records of Gastrotricha from South Africa, with description of a new species of Halichaetonotus (Chaetonotida, Chaetonotidae). ZooKeys, 142, 1–13. http://dx.doi.org/10.3897/zookeys.142.2036 Todaro, M.A., Dal Zotto, M., Jondelius, U., Hochberg, R., Hummon, W.D., Kanneby, T. & Rocha, C.E. (2012a) Gastrotricha: A Marine Sister for a Freshwater Puzzle. PLoS ONE 7, e31740. http://dx.doi.org/10.1371/journal.pone.0031740 Todaro, M.A., Guidi, L., Ferraguti, M. & Balsamo, M. (2012b) A fresh look at Dinodasys mirabilis (Gastrotricha, Macrodasyida), with focus on the reproductive apparatus and sperm ultrastructure. Zoomorphology, 131, 115–125. http://dx.doi.org/10.1007/s00435-012-0147-2 Todaro, M.A., Fleeger, J.W., Hu, Y.P., Hrincevich, A.W. & Foltz, D.W. (1996) Are meiofauna species cosmopolitan? Morphological and molecular analysis of Xenotrichula intermedia (Gastrotricha: Chaetonotida). Marine Biology, 125, 735–742. http://dx.doi.org/10.1007/BF00349256 Todaro, M.A. & Hummon, W.D. (2008) An overview and a dichotomous key to genera of the phylum Gastrotricha. Meiofauna Marina, 16, 3–20. Todaro, M.A. & Leasi, F. (2013) A new eye-bearing Macrodasys (Gastrotricha, Macrodasyida) from Jamaica. Meiofauna Marina, 20, 33–38. Todaro, M.A., Leasi, F. & Hochberg, R. (2014) A new species, genus and family of marine Gastrotricha from Jamaica, with a phylogenetic analysis of Macrodasyida based on molecular data. Systematics and Biodiversity, 12, 473–488. http://dx.doi.org/10.1080/14772000.2014.942718 Todaro, M.A., Perissinotto, R. & Bownes, S.J. (2013) Neogosseidae (Gastrotricha, Chaetonotida) from the iSimangaliso Wetland Park, KwaZulu-Natal, South Africa. ZooKeys, 315, 77–94. http://dx.doi.org/10.3897/zookeys.315.5593

208 · Zootaxa 3905 (2) © 2015 Magnolia Press TODARO ET AL.